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''Apoica flavissima'' is a paper wasp found primarily in South America. The species is distinguishable by its light coloring, unique single comb nests, and nocturnal nature. A notable feature of this species is the size dimorphism between queens and workers. Unlike most
Vespidae The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', '' Vespa orientalis'', and ''Vespula germanica'') and many solitary wasps. Eac ...
wasps, ''Apocia flavissima'' queens are smaller than their worker counterparts which results in unique intraspecies relationships.


Taxonomy and phylogeny

Originally, ''Apoica flavissima'' was thought to be a variety of ''
Apoica pallens The Central American paper wasp (''Apoica pallens'') is a nocturnality, nocturnal eusociality, eusocial wasp (family (biology), family Vespidae). It is famous for its swarm based emigration behavior, and is native to the lowlands of Central and ...
'' because of similar color and physical characteristics. It wasn't until 1972 that J. Van Der Vecht identified that three distinct species were mistakenly being categorized as one. Today, ''A. pallens'', ''A. flavissima'', and ''A. gelida'' are identified by differences in male genitalia. In addition, distinction can be made through slight color differences. While ''A. flavissima'' are entirely pale yellow, ''A. pallens'' are mostly yellow but have brown legs. Phylogenetic analysis indicates that ''Apoica flavissima'' is most closely related to ''Apoica pallens''.


Description

''A. flavissima'' typically have a mesoscutum (the middle
thoracic The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cr ...
segment of the insect) that is dark yellow with outstanding bristles. The gaster (the enlarged posterior of the abdomen) is pale yellow along with the humeral plate. The posterior
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-le ...
of the species are widely separated from the eyes. In addition, the wing length is typically . Colonies of ''A. flavissima'' can be identified morphologically by having smaller queens than workers. Unlike most wasp species, ''A flavissima'' exhibit a morphological caste difference in which the queen is smaller than workers in overall size and in other notable measurements. Specifically, workers have larger
alitrunk The mesosoma is the middle part of the body, or tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the metasoma. It bears the legs, and, in the case of winged insects, the wings. In hymenopterans of t ...
length and number of
hamuli A hamus or hamulus is a structure functioning as, or in the form of, hooks or hooklets. Etymology The terms are directly from Latin, in which ''hamus'' means "hook". The plural is ''hami''. ''Hamulus'' is the diminutive – hooklet or little ho ...
. In addition to being smaller, the queens also exhibit color differences from the workers. Queens typically have a darker brown
frons Insect morphology is the study and description of the physical form of insects. The terminology used to describe insects is similar to that used for other arthropods due to their shared evolutionary history. Three physical features separate ins ...
, clypeus, and mandible, while workers exhibit a light yellow coloring. Workers typically have a dark brown coloring of the basal half of the first gastral
tergite A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'mar ...
while queens are typically light yellow. The most striking morphological difference between queens and workers relates to the lateral tip of the
pronotum The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum ( dorsal), the prosternum (ventral), and the propleuron ( lateral) on ...
. Queens exhibit a tip that is gradually rounded while workers have a more acutely curved tip. Nests are also a defining feature as they only have one comb. Young nests tend to appear hexagonal and grow by curving downward and becoming increasingly rounded and oval. The texture of the nests are described as felt-like.


Distribution and habitat

''Apoica flavissma'' are located in South America. The species can be primarily found in
Panama Panama ( , ; es, link=no, Panamá ), officially the Republic of Panama ( es, República de Panamá), is a List of transcontinental countries#North America and South America, transcontinental country spanning the Central America, southern ...
,
Guyana Guyana ( or ), officially the Cooperative Republic of Guyana, is a country on the northern mainland of South America. Guyana is an indigenous word which means "Land of Many Waters". The capital city is Georgetown Guyana, Georgetown. Guyana ...
,
Suriname Suriname (; srn, Sranankondre or ), officially the Republic of Suriname ( nl, Republiek Suriname , srn, Ripolik fu Sranan), is a country on the northeastern Atlantic coast of South America. It is bordered by the Atlantic Ocean to the north ...
,
French Guiana French Guiana ( or ; french: link=no, Guyane ; gcr, label= French Guianese Creole, Lagwiyann ) is an overseas department/region and single territorial collectivity of France on the northern Atlantic coast of South America in the Guianas. It ...
,
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
,
Paraguay Paraguay (; ), officially the Republic of Paraguay ( es, República del Paraguay, links=no; gn, Tavakuairetã Paraguái, links=si), is a landlocked country in South America. It is bordered by Argentina to the south and southwest, Brazil to th ...
, and
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. This species is found in the
Neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bioge ...
region, which has some of the greatest biodiversity on earth. Nests are typically found in wooded areas that can provide shade to help with the
thermoregulation Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
of the nest. Due to the
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
nature of ''Apoica flavissima'', the overall abundance of the species is unknown. It is assumed that the species is relatively common.


Colony cycle

Colonies of ''Apoica flavissima'' are produced by the colony emigration of a swarm of at least one queen and multiple workers. ''A. flavissima'' differ from many other wasp species during this emigration because some males swarm along with females during this flight. Once a colony is founded, the cycle of an ''Apoica flavissima'' colony can be broken into three phases. The first phase is the pre-emergence phase. In this phase, no adult offspring have been produced, only eggs and young larvae. In the second stage, female workers are produced. The third stage is characterized by the production of males and queens. ''Apoica flavissima'' exhibits pre-imaginal determination of caste, meaning that an individual's caste is determined in the larval stage.


Behavior


Nocturnal nature

Observations of ''A. flavissima'' indicate that adult wasps are dormant without any visible movement during the daytime. The only exception is when the colony is disturbed and swarms in a communal defense of the nest. While in this dormant state, the individuals in the colony rest on the under surface of the comb. At nightfall, ''A. flavissima'' begin to abruptly depart the nest in an explosive swarm. Frequencies of departure appear to be affected by environmental factors such as cloudiness or rain. In addition, the foraging activities of ''A. flavissima'' correspond with phases of the moon as frequency of foraging significantly increases during the full and last quarter moon. This indicates that the presence of light is an important determinant of the nocturnal activities of the species.


Thermoregulation of nests

Since ''A. flavissma'' are observed to be dormant during the day, the species appears to have developed a behavior to regulate the temperature of the nest. This is done by coating the roof of the nest with oral secretions and attaching plant fibers to the roof of the nest. The central part of the roof is made to be particularly thick, to help with the insulation of the structure. This technique is successful at maintaining a mean temperature of 27.3 °C in the nest. During the day when temperature fluctuations are greater, the generation of metabolic heat from individuals in the nest appears to be key to the temperature stabilization of the nest.


Kin selection


Pre-determination of caste

While some species of wasp select their queen on the basis of size, the selection of ''A. flavissima'' queens is physically pre-determined. Workers have under developed ovaries and are incapable of producing offspring throughout their entire lifestyle. Queens are the only females with developed ovaries and have long ovarioles with two or three mature oocytes. In contrast, worker ''A. flavissima'' exhibit underdeveloped ovaries with no visible oocytes. This physical morphological caste difference means that the ability of an individual to reproduce does not change in respect to environmental constraints or in respect to the life stage of the nest. This means there are no intermediates between queens and workers, creating a strict social hierarchy within the nest.


Genetic relatedness within colonies

Colonies of ''A. flavissima'' can rotate between polygyny (multiple queens) and monogyny (one queen). Due to this, the relatedness of workers to the queen or queens varies through time. Large nests typically have more queens, so relatedness between individuals could correlate with the size of the nest. Since it is often believed that workers rear the young of the queen altruistically due to relatedness of females, this fluctuation may lead to conflicts between castes in large nests. Due to this, in cases where the nest has multiple queens, these queens have little control over their reproductive outputs. Accordingly, as few as 1.5% of queens may be functional within a nest.


Worker-queen conflict

In ''Apoica flavissima'' nests, workers appear to have behavioral control. This may be caused by the low relatedness between females or the distinct morphological caste differences. Workers have been observed to behaviorally police the queen's reproductive output by biting and harassment. In large nests with multiple queens, workers have even been known to remove queens from the nest. These conflicts indicate that the workers play a part in the reproductive output of the nest.


Interaction with other species


Diet

''Apoica flavissima'' is observed to survive primarily on nectar brought back to the nest to feed young and queens during night foraging. Most of the materials brought back to the nest are given to a small collection of nest-mates who are tasked with distributing the food to feed both larvae and other adults. Recent studies have indicated that ''Apoica flavissima'' may also exhibit necrophagy, or the consumption of flesh, on larger vertebrates and invertebrate carcasses. Evidence of this behavior is a structural modification of the
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
of the species. ''A. flavissima'' has a dorsal tooth modification of the inner surface of its mandibles where the tooth is elongated and blade-like, making it potentially possible for the species to consume the flesh of bigger organisms.


Parasitism

''A. flavissima'' are occasionally subjected to parasitism by other wasp species. Wasps in the family
Trigonalidae Trigonalidae (the spelling Trigonalyidae is incorrect under Article 29.5 of the ICZN,ICZN Code
29.5. Mainten ...
are known for their parasitic nature, relying on hosts to ingest their eggs or provide homes for their young. ''A. flavissima'' appear to be subject to the latter, acting as a secondary host to trigonalid species who invade the nest and grow their young. Recent research indicates that some species of trigonalids may use ''A. flavissima'' as a primary host, using the species to inject their larvae and transfer it to the nest via
trophallaxis Trophallaxis () is the transfer of food or other fluids among members of a community through mouth-to-mouth ( stomodeal) or anus-to-mouth ( proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of molecules such as pher ...
.


References

{{Taxonbar, from=Q14423246 Hymenoptera of South America Vespidae Insects described in 1972